Do group dynamics affect colour morph clines during a range shift?

Species exhibiting colour polymorphism are thought to have an ecological advantage at the landscape scale, because spatial segregation of alternatively adapted ecotypes into diverse habitats can increase the species' niche breadth and thus confer greater geographic range size. However, morph frequencies are also influenced by intrapopulational processes such as frequency- or density-dependent social interactions. To identify how social feedback may affect clinal variation in morph frequencies, we investigated reciprocal interactions between morph-specific thermal tolerance, local climatic conditions and social environments, in the context of a colour-morph frequency cline associated with a recent range expansion in blue-tailed damselflies (Ischnura elegans) in Sweden. Cold tolerances of gynochromes (female-like female morph) were positively correlated with local gynochrome frequencies, suggesting a positive frequency-dependent fitness benefit. In contrast, androchrome (male-mimic female morph) cold tolerances were improved following recent exposure to cold weather, suggesting a beneficial environmental acclimation effect. Thus, according to an environment-matching hypothesis for clinal variation, androchrome frequencies should therefore increase towards the (cooler) range limit. In contrast to this prediction, gynochrome frequencies increased at the expanding range limit, consistent with a positive frequency-dependent social feedback that is beneficial when invading novel climates. Our results suggest that when phenotypes or fitnesses are affected by interactions with conspecifics, beneficial social effects on environmental tolerances may (i) facilitate range shifts, and (ii) reverse or counteract typical patterns of intraspecific interactions and environment-matching clines observed in stable populations observed over broader geographic scales.

Rapid increase in dispersal during range expansion in the invasive ladybird Harmonia axyridis.

The evolutionary trajectories associated with demographic, genetic and spatial disequilibrium have become an issue of growing interest in population biology. Invasive species provide unique opportunities to explore the impact of recent range expansion on life-history traits, making it possible to test for a spatial arrangement of dispersal abilities along the expanding range, in particular. We carried out controlled experiments in laboratory conditions to test the hypothesis of an increase in dispersal capacity with range expansion in Harmonia axyridis, a ladybird that has been invading Europe since 2001. We found a marked increase in the flight speed of the insects from the core to the front of the invasion range in two independent sampling transects. By contrast, we found that two other traits associated with dispersal (endurance and motivation to fly off) did not follow the same spatial gradient. Our results provide a striking illustration of the way in which predictable directional genetic changes may occur rapidly for some traits associated with dispersal during biological invasions. We discuss the consequences of our results for invasion dynamics and the evolutionary outcomes of spatially expanding populations.

Sex differences in morphology across an expanding range edge in the flightless ground beetle, Carabus hortensis.

Species' ranges are dynamic, changing through range shifts, contractions, and expansions. Individuals at the edge of a species' shifting range often possess morphological traits that increase movement capacity, that are not observed in individuals farther back within the species' range. Although morphological traits that increase in proportion toward the range edge may differ between the sexes, such sex differences are rarely studied.Here, we test the hypotheses that body size and condition increase with proximity to an expanding range edge in the flightless ground beetle, Carabus hortensis, and that these trait changes differ between the sexes.Male, but not female, body size increased with proximity to the range edge. Body size was positively correlated with male front and mid tibia length and to female hind tibia length, indicating that body size is indicative of movement capacity in both sexes. Body condition (relative to body size) decreased with increasing population density in males but not females. Population density was lowest at the range edge.Our results indicate that sex is an important factor influencing patterns in trait distribution across species' ranges, and future studies should investigate changes in morphological traits across expanding range margins separately for males and females. We discuss the implications for sex differences in resource allocation and reproductive rates for trait differentiation across species' shifting ranges.

Invasion Dynamics of A Termite, Reticulitermes flavipes, at Different Spatial Scales in France.

Termites are social insects that can also be major pests. A well-known problem species is the subterranean termite, Reticulitermes flavipes. It is invasive in France and is thought to have arrived from Louisiana during the 18th century. While the putative source of French populations has been identified, little is known about how the termite spread following its establishment. Here, we examined expansion patterns at different spatial scales in urban areas to clarify how R. flavipes spread in France. Based on our analyses of phylogeography and population genetics, results suggest a scenario of successive introductions into the Charente-Maritime region, on the Atlantic Coast. Two major expansion fronts formed: one that spread toward the northeast and the other toward the southeast. At the regional scale, different spatial and genetic distribution patterns were observed: there was heterogeneity in Île-de-France and aggregation in Centre-Val de Loire. At the local scale, we found that our three focal urban sites each formed a single large colony that contained several secondary reproductives. Our findings represent a second step in efforts to reconstruct termite's invasion dynamics. They also highlight the role that may have been played by the French railway network in transporting termites over long distances.